采后BTH處理對(duì)蘋果果實(shí)苯丙烷代謝和病程相關(guān)蛋白積累的增強(qiáng)作用

葛永紅,李燦嬰,朱丹實(shí),董柏余,蘭 琳

(1.渤海大學(xué)化學(xué)化工與食品安全學(xué)院,遼寧錦州 121013;2.甘肅農(nóng)業(yè)大學(xué)食品科學(xué)與工程學(xué)院,甘肅蘭州 730070)

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采后BTH處理對(duì)蘋果果實(shí)苯丙烷代謝和病程相關(guān)蛋白積累的增強(qiáng)作用

葛永紅1,李燦嬰1,朱丹實(shí)1,董柏余2,蘭 琳2

(1.渤海大學(xué)化學(xué)化工與食品安全學(xué)院,遼寧錦州 121013;2.甘肅農(nóng)業(yè)大學(xué)食品科學(xué)與工程學(xué)院,甘肅蘭州 730070)

本文探討采后苯丙噻重氮(BTH)處理對(duì)蘋果果實(shí)苯丙烷代謝和病程相關(guān)蛋白產(chǎn)生的增強(qiáng)作用。以“國光”蘋果為試材,采后用100mg/L BTH浸泡處理10min,測(cè)定處理對(duì)果實(shí)苯丙烷代謝相關(guān)酶活性、產(chǎn)物積累和病程相關(guān)蛋白產(chǎn)生的影響。結(jié)果表明,BTH處理顯著提高了蘋果果實(shí)體內(nèi)苯丙氨酸解氨酶(PAL)、4-香豆素輔酶A連接酶(4CL)的活性,增加了總酚、類黃酮及木質(zhì)素含量,同時(shí)提高了β-1,3葡聚糖酶(GLU)、幾丁質(zhì)酶(CHT)和過氧化物酶(POD)的活性。由此表明,采后BTH處理可通過誘導(dǎo)果實(shí)苯丙烷代謝和促進(jìn)病程相關(guān)蛋白的產(chǎn)生來增強(qiáng)果實(shí)對(duì)采后病害的抗性。

果實(shí),BTH,苯丙烷代謝,病程相關(guān)蛋白,誘導(dǎo)抗性

蘋果(MalusdomesticaBorkh)屬于薔薇科(Rosaceae)蘋果屬(Malus)植物,是典型的呼吸躍變型果實(shí),在采后貯藏、運(yùn)輸和銷售過程中很容易腐爛變質(zhì),給生產(chǎn)者和經(jīng)營者帶來巨大的經(jīng)濟(jì)損失。由病原物引起的采后病害是導(dǎo)致果蔬爛損的重要原因,每年的損失在發(fā)展中國家可占其果蔬總產(chǎn)量的30%～50%,在發(fā)達(dá)國家可占10%～30%[1]。由Penicilliumexpansum引起的青霉病是蘋果采后的主要病害之一[2],傳統(tǒng)的控制方式是采用化學(xué)殺菌劑如噻菌靈、甲基硫菌靈、嘧霉胺和撲海因等[3-4],但存在殺菌劑殘留、環(huán)境污染和病原物產(chǎn)生抗藥性等問題[5]。誘導(dǎo)果蔬的抗病性已成為近年來控制或減輕采后病害的新手段[3-5]。苯丙噻重氮(benzothiadiazole,BTH)屬水楊酸類似物,是第一個(gè)人工合成的化學(xué)誘抗劑。在有效濃度范圍內(nèi),BTH不表現(xiàn)直接的殺菌活性,但能誘導(dǎo)植物產(chǎn)生抗病性,并且不會(huì)對(duì)環(huán)境造成污染[6]。已有研究報(bào)道BTH處理能夠誘導(dǎo)多種果實(shí)的抗病性如草莓、桃、梨、蘋果、厚皮甜瓜、芒果、馬鈴薯等[7-14]。BTH誘導(dǎo)果實(shí)產(chǎn)生抗病性的機(jī)理主要涉及活性氧的產(chǎn)生、活化苯丙烷代謝途徑、積累病程相關(guān)蛋白等[15]。我們的前期研究發(fā)現(xiàn)采后BTH處理能夠顯著降低蘋果果實(shí)的青霉病,并且誘導(dǎo)了果實(shí)體內(nèi)活性氧的產(chǎn)生及活性氧代謝相關(guān)酶活性的提高,但關(guān)于其他的機(jī)理沒有進(jìn)行系統(tǒng)的研究。

本實(shí)驗(yàn)以蘋果為試材,在前期研究的基礎(chǔ)上,重點(diǎn)探討采后BTH處理對(duì)果實(shí)苯丙烷代謝相關(guān)酶苯丙氨酸解氨酶(PAL)、4-香豆素輔酶A連接酶(4CL)及主要產(chǎn)物總酚、類黃酮、木質(zhì)素等的積累和β-1,3葡聚糖酶(GLU)、幾丁質(zhì)酶(CHT)和過氧化物酶(POD)等病程相關(guān)蛋白產(chǎn)生的影響,為闡明BTH誘導(dǎo)蘋果抗病性提供理論依據(jù)。

1 材料與方法

1.1 材料與儀器

國光蘋果于2013年9月采自甘肅省國營條山農(nóng)場(chǎng),單果包裝入紙箱后當(dāng)天運(yùn)抵甘肅農(nóng)業(yè)大學(xué)采后生物學(xué)實(shí)驗(yàn)室,常溫((22±2)℃,RH 50%～60%)下貯藏待用;BTH由Novatis公司提供(有效濃度50%)。

UV-2450型分光光度計(jì) 日本島津;H-1850R型離心機(jī) 長沙湘儀離心機(jī)儀器有限公司;WYX-A微型旋渦混合器 上海躍進(jìn)醫(yī)療器械廠。

1.2 實(shí)驗(yàn)方法

1.2.1 BTH處理 大小均勻,無機(jī)械損傷和病蟲害的果實(shí)分別用100mg/L BTH溶液和清水浸泡(對(duì)照)10min(內(nèi)含0.05% Tween 20),取出晾干后入包裝箱,于常溫條件下((22±2)℃,RH 55%～60%)貯藏待用,每處理用果實(shí)30個(gè),重復(fù)3次。

1.2.2 取樣 參照范存斐等方法[16]。BTH和清水處理后的果實(shí)分別于0、2、4、6、8d取皮下1～5mm 處果肉組織3g,鋁箔紙包裹,用液氮速凍后在-80℃超低溫冰箱中保存待用。

1.2.3 苯丙氨酸解氨酶(PAL)活性測(cè)定 參照Liu等方法[8]。酶反應(yīng)體系包括:2mL 0.2mol/L硼酸緩沖液(pH8.8),300μL酶粗提液和1mL L-苯丙氨酸(0.02mol/L)。24℃下反應(yīng)2min 后測(cè)其OD290?？瞻滓跃彌_液代替酶粗提液。以U/g FW(鮮重)表示酶活性,每分鐘內(nèi)OD290變化0.01為1個(gè)活性單位(U)。

1.2.4 4-香豆素-輔酶A連接酶(4 CL)活性測(cè)定 參照范存斐等方法[16]。以每分鐘吸光值變化0.01為1個(gè)活性單位(U),4 CL活性以U/g FW表示。

1.2.5 總酚和類黃酮含量的測(cè)定 參照范存斐等方法[16]?？偡雍恳設(shè)D280/g FW表示,類黃酮含量表示為OD325/g FW。

1.2.6 木質(zhì)素含量的測(cè)定 參照范存斐等方法[16]。木質(zhì)素含量以O(shè)D280/g FW表示。

1.2.7 過氧化物酶(POD)活性的測(cè)定 參照葛永紅等方法[17]并修改。酶反應(yīng)體系包括1.8mL愈創(chuàng)木酚(0.05mol/L),30μL酶粗提液和100μL,2% H2O2,然后于470nm波長下測(cè)定吸光度的變化,以每分鐘內(nèi)OD470變化0. 01為1個(gè)活性單位(U),POD活性以U/g FW表示。

1.2.8 β-1,3-葡聚糖酶(GLU)活性的測(cè)定 參照Ge等方法[18]并修改。酶反應(yīng)體系包括0.8mL,1mg/mL昆布多糖,0.2mL酶液,2mL銅試劑,2mL砷鉬酸試劑,反應(yīng)后在660nm處測(cè)定吸光度,對(duì)照標(biāo)準(zhǔn)曲線求出樣品液中的葡萄糖量。在最適反應(yīng)條件下(37℃)下每秒鐘每升酶液催化底物昆布多糖產(chǎn)生1μmol的葡萄糖為一個(gè)酶活力單位(U)。

1.2.9 幾丁質(zhì)酶(CHT)活性的測(cè)定 參照葛永紅等方法[17]。以每分鐘生成1μmol N-乙酰氨基葡萄糖所需的酶量為一個(gè)酶活力單位(U)。

1. 3 數(shù)據(jù)處理

全部實(shí)驗(yàn)數(shù)據(jù)用Microsoft Excel 2003 處理并作圖,用SPSS16.0統(tǒng)計(jì)軟件進(jìn)行LSD統(tǒng)計(jì)分析,圖中以標(biāo)準(zhǔn)誤差(±SE)表示。

2 結(jié)果與分析

2.1 BTH處理對(duì)蘋果果實(shí)PAL、4CL活性的影響

PAL 是苯丙烷代謝途徑的一個(gè)限速酶,其活性高低與酚類、黃酮類和木質(zhì)素的積累密切相關(guān)。隨著貯藏時(shí)間的延長,蘋果果實(shí)PAL先升高后降低,但BTH處理顯著提高了果實(shí)PAL活性,在貯藏第4d達(dá)到最大,高出對(duì)照71.4%,第8d BTH處理果實(shí)PAL活性又略有升高,而對(duì)照則呈下降趨勢(shì)(圖1A)。4CL活性的變化趨勢(shì)基本和PAL一致,BTH處理也顯著提高了蘋果果實(shí)4CL活性,在貯藏第6d活性達(dá)到最大,高出對(duì)照32.2%(圖1B)。

圖1 采后BTH處理對(duì)蘋果果實(shí)PAL(A) 和4CL(B)活性的影響Fig.1 Effect of postharvest BTH dipping treatment on PAL(A)and 4CL(B)activities in apple fruit

2.2 BTH處理對(duì)蘋果果實(shí)總酚和類黃酮含量的影響

總酚和類黃酮是苯丙烷代謝的產(chǎn)物,其含量高低與果實(shí)預(yù)防病原菌的侵入有關(guān)。處理和對(duì)照蘋果果實(shí)總酚含量在貯藏期間總體呈先升高后下降趨勢(shì),但BTH處理顯著提高了果實(shí)總酚含量,并且在貯藏第4d出現(xiàn)高峰,高出對(duì)照25.9%(圖2A)。貯藏期間果實(shí)類黃酮含量先升高后降低,BTH處理果實(shí)類黃酮含量顯著高于對(duì)照果實(shí),如在貯藏第4d時(shí)類黃酮含量高出對(duì)照27.2%(圖2B)。

圖2 采后BTH處理對(duì)蘋果果實(shí)總酚(A) 和類黃酮(B)含量的影響Fig.2 Effect of postharvest BTH dipping treatment on the content of total phenolic compounds(A) and flavonoids(B)in apple fruit

2.3 BTH處理對(duì)蘋果果實(shí)木質(zhì)素含量的影響

木質(zhì)素是苯丙烷代謝途徑的產(chǎn)物,果實(shí)中木質(zhì)素的積累有利于增強(qiáng)細(xì)胞壁厚度,從而提高對(duì)病原物的抵抗力。蘋果果實(shí)在貯藏期間,木質(zhì)素含量總體呈先升高后下降趨勢(shì),對(duì)照果實(shí)中木質(zhì)素含量一直維持在一個(gè)較低的水平,但BTH處理果實(shí)中木質(zhì)素含量顯著高于對(duì)照,并且在貯藏第6d達(dá)到最大,高出對(duì)照35.4%(圖3)。

圖3 采后BTH處理對(duì)蘋果果實(shí)木質(zhì)素含量的影響Fig.3 Effect of postharvest BTH dipping treatment on lignin content in apple fruit

2.4 BTH處理對(duì)蘋果果實(shí)POD活性的影響

POD廣泛存在于植物中,在果實(shí)體內(nèi)主要參與活性氧的清除、木質(zhì)素的合成等過程。在整個(gè)貯藏期間對(duì)照和BTH處理果實(shí)POD活性都呈先上升后下降的趨勢(shì),但是BTH處理果實(shí)POD活性顯著高于對(duì)照,如在貯藏第4、6d分別高出對(duì)照49.8%和55.8%(圖4)。

圖4 采后BTH處理后蘋果果實(shí)POD活性的變化Fig.4 Changes in the activity of POD in apple fruit after BTH dipping treatment

2.5 BTH處理對(duì)蘋果果實(shí)GLU和CHT活性的影響

GLU和CHT是兩種主要的病程相關(guān)蛋白,主要用來降解病原物細(xì)胞壁,從而導(dǎo)致病原物死亡。貯藏期間對(duì)照果實(shí)GLU活性總體維持在一定水平,而BTH處理顯著提高果實(shí)中GLU活性,在貯藏第6d出現(xiàn)活性高峰,高出對(duì)照63.7%(圖5A)。CHT活性總體呈先升高后降低趨勢(shì),但BTH處理果實(shí)中CHT活性顯著高于對(duì)照果實(shí),在貯藏第6d達(dá)到最大,高出對(duì)照30.3%(圖5B)。

圖5 采后BTH處理后蘋果果實(shí)GLU(A) 和CHT(B)活性的變化Fig.5 Changes in the activity of GLU(A)and CHT(B) in apple fruit after BTH dipping treatment

3 討論

苯丙烷代謝是植物和果蔬中次生代謝產(chǎn)物合成的主要途徑,在抗病中具有重要的作用[19]。PAL和4CL是該途徑中的關(guān)鍵酶[25],PAL能夠調(diào)控植物和果實(shí)中酚類、黃酮類、生物堿、花青素、木質(zhì)素等抗菌物質(zhì)的合成,木質(zhì)素的合成有利于強(qiáng)化細(xì)胞壁,增強(qiáng)對(duì)病原物侵染的抵抗力[20],而4CL主要調(diào)控苯丙烷代謝主途徑向分支途徑的轉(zhuǎn)折[21]。本實(shí)驗(yàn)表明,BTH能顯著誘導(dǎo)蘋果果實(shí)中PAL和4CL活性的升高,同時(shí)促進(jìn)了果實(shí)中總酚、類黃酮和木質(zhì)素的積累。桃[8]、梨[9],厚皮甜瓜[22]、草莓[23]等果實(shí)的研究中也得到了類似的結(jié)果。本實(shí)驗(yàn)還發(fā)現(xiàn)BTH處理蘋果果實(shí)貯藏后期PAL活性有所升高,這與我們?cè)谔鸸瞎麑?shí)上的研究結(jié)果類似[22],可能是因?yàn)榍捌赑AL和4CL活性的升高促進(jìn)了黃酮類、生物堿和花青素的積累,而木質(zhì)素的合成過程比較復(fù)雜,需要在POD、過氧化氫及漆酶的共同作用下合成,因此后期PAL的升高主要是調(diào)控木質(zhì)素的合成。還有研究發(fā)現(xiàn)采前BTH處理也能誘導(dǎo)甜瓜果實(shí)中苯丙烷代謝產(chǎn)物的積累,從而降低潛伏侵染的發(fā)生率[28]。酚類物質(zhì)在多酚氧化酶(PPO)的作用下氧化成對(duì)病原菌產(chǎn)生高毒性的醌類物質(zhì),直接抑制侵染寄主的病原物生長[24]。此外,酚類物質(zhì)還可以作為木質(zhì)素合成的前體物質(zhì),促進(jìn)細(xì)胞壁中木質(zhì)素的積累,從而增厚細(xì)胞壁提高對(duì)病原物抵抗力[25-26]。類黃酮具有直接的抑菌活性,可抑制真菌孢子萌發(fā)、芽管伸長和菌絲生長[27]。由此表明,采后BTH處理可明顯激活蘋果果實(shí)體內(nèi)酚類和類黃酮合成途徑,以增加果實(shí)對(duì)病原物侵染的抵抗能力。

病程相關(guān)蛋白(Pathogenesis-related proteins,PRs)是植物受到病原菌侵染或非生物脅迫條件下產(chǎn)生的小分子蛋白質(zhì),在抗病過程中發(fā)揮著重要作用[21]。目前已經(jīng)發(fā)現(xiàn)的PRs有17種,根據(jù)其結(jié)構(gòu)和功能的不同表示為PR-1至PR-17,其中PR-2和PR-3,8,11分別具有GLU和CHT活性,可以直接分解真菌細(xì)胞壁主要成分為葡聚糖和幾丁質(zhì),破壞其結(jié)構(gòu)[21]。本研究結(jié)果發(fā)現(xiàn),BTH處理可顯著提高GLU和CHT活性。已有研究報(bào)道,采前或采后BTH處理能夠誘導(dǎo)甜瓜[28]、桃[8,29]、芒果[30]、梨[31]果實(shí)中GLU和CHT的活性,從而提高果實(shí)的抗病性。PR9具有POD活性,參與果實(shí)體內(nèi)木質(zhì)素的合成、細(xì)胞壁結(jié)構(gòu)蛋白的交聯(lián)以及活性氧的清除等[15,21]。本研究結(jié)果表明,BTH處理誘導(dǎo)了POD活性的提高,促進(jìn)了木質(zhì)素含量的積累,提高了果實(shí)的抗病性。這與在甜瓜[17]、砂糖橘[32]、芒果[33]上的研究結(jié)果相一致。此外,BTH處理還能誘導(dǎo)芒果和番木瓜果實(shí)的過氧化物酶基因(POD)表達(dá)[34-35]。有關(guān)BTH處理后蘋果果實(shí)中苯丙烷代謝和病程相關(guān)蛋白基因表達(dá)有待進(jìn)一步分析研究。

4 結(jié)論

采后100mg/L BTH處理顯著提高了蘋果果實(shí)PAL和4CL活性,并且促進(jìn)了苯丙烷代謝產(chǎn)物總酚、類黃酮和木質(zhì)素含量的提高。BTH處理還促進(jìn)了蘋果果實(shí)POD,GLU和CHT活性的提高。由此表明,采后BTH處理可通過激活蘋果果實(shí)苯丙烷代謝途徑增強(qiáng)組織結(jié)構(gòu),促進(jìn)病程相關(guān)蛋白的產(chǎn)生,從而提高了果實(shí)抵抗病原物侵染的能力。

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Activation of postharvest BTH treatment on phenylpropanoid pathwayand pathogenesis-related proteins accumulation in apple fruit

GE Yong-hong1,LI Can-ying1,ZHU Dan-shi1,DONG Bo-yu2,LAN Lin2

(1.College of Chemistry,Chemical Engineering and Food Safety,Jinzhou 121013,China;2.College of Food Science and Engineering,Gansu Agricultural University,Lanzhou 730070,China)

The effect of benzothiadiazole(BTH)dipping on phenylpropanoid pathway and accumulation of pathogenesis-related proteins in apple fruit was investigated in this paper. Apple fruit(cv. Guoguang)were dipped in 100mg/L BTH for 10min to study the effect of BTH on key enzymes activities,metabolites accumulation in phenylpropanoid pathway and production of pathogenesis-related proteins(PRs). The results indicated that BTH treatment significantly increased the activities of phenylalanine ammonia-lyase(PAL)and 4-coumaric acid-coenzyme A ligase(4CL),and the content of total phenolic compounds,flavonoids and lignin. The activities of β-1,3-glucanase(GLU),chitinase(CHT)and peroxidase(POD)were also induced by BTH treatment. These results suggest that postharvest BTH treatment induced disease resistance in apple fruit by activating phenylpropanoid pathway and promoted the accumulation of PRs.

fruit;benzothiadiazole(BTH);phenylpropanoid pathway;pathogenesis-related proteins;induced resistance

2014-06-03

葛永紅(1979-)，男，博士，副教授，研究方向：果蔬貯藏與保鮮。

國家自然科學(xué)基金(31160405)；渤海大學(xué)博士啟動(dòng)基金(bsqd201405)；遼寧省食品安全重點(diǎn)實(shí)驗(yàn)室項(xiàng)目(LNSAKF2013021)。

TS255.1

A

1002-0306(2015)05-0306-05

10.13386/j.issn1002-0306.2015.05.056